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tuls ennn 

i LompaFadfe JL 




US ISSN 0006-9698 

Cambridge, Mass. 

8 May 2012 

Number 529 



Juan D. Daza 1 and Aaron M. Bauer 1 - 2 

Abstract. A new species of Sphaerodactylus (Squamata: Gekkota: Sphaerodactylidae) is described from an 
amber inclusion from the late Early Miocene or early Middle Miocene (15 to 20 million years ago) of the Dominican 
Republic. Unlike earlier amber-embedded specimens assigned to this genus, the new specimen is largely skeletal, with 
some integument remaining. A combination of 258 (of 674) osteological and external characters could be scored for 
the new species in a cladistic analysis of 21 gekkotan species, including representatives of all sphaerodactylid genera. 

The most parsimonious trees obtained confirm the placement of the amber gecko within the genus Sphaerodactylus 
and a comparison with extant Hispaniolan and Puerto Rican congeners suggests phenetic similarity both with 
members of S. difficilis complex and the S. shrevei species group. Character mapping on the basis of the phylogenetic 
analysis permits the preliminary identification of morphological characters diagnostic of the Sphaerodactylidae, 
Sphaerodactylini, and Sphaerodactylus. Osteological features of the new species are discussed in the broader context 
of sphaerodactyl, sphaerodactylid, and gekkotan variation. Extant Hispaniolan Sphaerodactylus display significant 
ecomorphological variation and it is likely that the many known, though not yet described, amber-embedded 
specimens will eventually reveal similar patterns in their Miocene congeners. 

Key words: Sphaerodactylus; Sphaerodactylidae; gecko; osteology; Hispaniola; new species; phylogeny 


Amber-embedded fossils provide unique 
insights into extinct vertebrate taxa as they 

1 Department of Biology, Villanova University, 800 Lancaster 
Avenue, Villanova, Pennsylvania 19085, U.S.A.; e-mail:, 

2 Department of Herpetology, Museum of Comparative 
Zoology, Harvard University, 26 Oxford Street, Cam- 
bridge, Massachusetts 02138, U.S.A. 

frequently preserve the integument and 
thus give an impression of what the intact 
animal looked like in life. The mode of 
amber preservation, however, limits verte- 
brate inclusions to taxa small enough to be 
trapped in the viscous resin. Thus, aside 
from isolated bird feathers (e.g., Grimaldi 
and Case, 1995; Alonso et al, 2000; 
Perrichot et al, 2008) and mammal hairs 
and bones (MacPhee and Grimaldi, 1996; 

The President and Fellows of Harvard College 2012. 


No. 529 

Sontag, 2008), known vertebrate inclusions 
are limited to several minute frogs (Poinar 
and Cannatella, 1987; Poinar, 1992; Grimaldi, 
1996), unidentified (but probably squamate) 
skin pieces (Poinar and Poinar, 1999; Grimaldi 
et al, 2002; Perrichot and Neraudeau, 2005), 
and to a fairly large number of small 
lizards. At least four areas of the world 
have yielded such lizard fossils. The oldest 
inclusion is Baabdasaurus xenurus, an au- 
tarchoglossan of indeterminate affinities 
described from a partial specimen in amber 
from the Lower Cretaceous (120 million 
years ago [MYA]) of Lebanon (Arnold 
et al, 2002). A 100-million-year-old gekko- 
tan in amber, Cretaceogekko burmae, is 
known from deposits in Myanmar (Arnold 
and Poinar, 2008). Amber deposits of the 
Baltic, dating from the early Eocene (Lars- 
son, 1978; Ritzkowski, 1997; Weitschat and 
Wichard, 2002), have yielded a minimum 
of three different species in the extinct 
lacertid genus Succinilacerta (Katinas, 1983; 
Kosmowska-Ceranowicz et al. , 1997a, 19976; 
Krumbiegel, 1998; Bdhme & Weitschat, 1998, 
2002; Borsuk-Biarynicka et al, 1999) and a 
single gekkotan, Yantarogekko balticus (Bauer 
etal, 2005). 

The greatest number of amber lizards, 
however, are known from the Miocene of the 
Dominican Republic. These fossils include 
several specimens referred to the dactyloid 
genus Anolis (Lazell, 1965; Rieppel, 1980; de 
Queiroz et al, 1998; Polcyn et al, 2002), and 
several geckos referable to the extant genus 
Sphaerodactylus (Bohme, 1984; Kluge, 1995). 
Although a very limited number of lizards in 
Dominican amber have been formally de- 
scribed, many more specimens are known to 
exist in the holdings of private collectors 
(Poinar and Poinar, 1999; D. A. Grimaldi, 
personal communication). 

Biostratigraphic and paleogeographic data 
indicate that the amberiferous deposits in the 
Dominican Republic were formed in a single 

sedimentary basin during the late Early 
Miocene through early Middle Miocene, 
about 15 to 20 MYA (Grimaldi, 1995; 
Iturralde-Vinent and MacPhee, 1996). Am- 
ber fossils from tropical America originate 
mainly from the resin of the extinct legumi- 
nous tree Hymenaea protera (Poinar and 
Cannatella, 1987; Poinar, 1992; Iturralde- 
Vinent, 2001), and on the basis of inference 
from historical forest distribution in Hispa- 
niola, these trees were mainly distributed in 
the evergreen forests of the southeast area, 
surrounding the depositional basin (Itur- 
ralde-Vinent and MacPhee, 1996). Today, 
Dominican amber is commercially exploited 
in three geological formations: La Toca 
(North), Yanigua (Eastern), and Sombrerito 
(south of the Cordillera Central in the area 
of Plateau Central-San Juan). 

We here report on a new amber-embedded 
gecko from La Toca mine, in the Cordillera 
Septentrional, Santiago Province, north of 
Municipio Santiago de los Caballeros 
(Fig. 1). We further review osteological data 
for the Sphaerodactylidae and identify puta- 
tive synapomorphies that support the mono- 
phyly of this recently recognized clade of 
gekkotans. To date only two nonmolecular 
characters, neither found in all members of 
the clade, have been identified as possible 
evidence of affinities: 

1 . presence of parafrontal bones (present 
in Aristelliger and Teratoscincus and pre- 
sumably lost in all miniaturized sphaerodac- 
tylids) and 

2. single-egg clutch (except in Teratoscincus 
and Euleptes, which retain two-egg clutches; 
Gamble et al, 2008a). 


Anatomical Observations. Faked lizard 
inclusions in amber are common, so the 
authenticity of the specimen was confirmed 
on the basis of several criteria intrinsic to both 



La Toca Formation 
Northern mining district 


Figure 1. Left, black areas indicate the distribution of Sphaerodactylus geckos in the Americas. Right, map of 
Hispaniola showing the type locality of Sphaerodactylus ciguapa sp. nov. (modified from Iturralde-Vinent, 2001). 

the amber itself and the lizard it contained 
(Bauer and Branch, 1995). The amber specimen 
and comparative ethanol-preserved, cleared 
and stained, and dry skeletal material were 
examined using a Nikon SMZ1000 dissecting 
microscope equiped with a digital camera 
(Nikon DS-Fil) and the image acquisition 
software NIS Elements D v. 3.1, and a Leica 
MZ6 dissecting microscope equipped with a 
camera lucida. Digital radiographs were ob- 
tained using a Kevex™ PXS10-16W X-ray 
source and Varian Amorphous Silicon Digital 
X-Ray Detector PaxScan® 4030R set to 130 kV 
at 8 1 uA. For each X-ray linear and pseudofilm 
filters were used. Drawings were traced directly 
over digital images using Adobe® Illustrator® 
CS3 13.0.2 and complemented with illustra- 
tions made with the camera lucida. Measure- 
ments were made from the X-ray images to 
avoid measurement error caused by the refrac- 
tive index of amber in = 1.55). Anatomical 
terminology follows Daza et al. (2008). 

Phylogenetic Analysis. To objectively iden- 
tify characters uniting the new species with its 
congeners and other members of the Spha- 
erodactylidae, a cladistic analysis of selected 
taxa was performed. A morphological data 

set of 674 characters scored for 21 taxa 
(Appendix 1 ; data available at www.morpho-; project number p532, accession 
number XI 201) was analyzed in the computer 
program T.N.T. (Goloboff et al, 2003a, 
2008) using maximum parsimony. All char- 
acters were treated as unordered, and equally 
weighted. In addition to representatives of 12 
sphaerodactylid genera, including all six 
genera of "sphaerodactyls," seven outgroup 
gekkotan species were included: Hemidactylus 
brookii, Narudasia festiva, and Pseudogekko 
smaragdinus (Gekkonidae), and Phyllodac- 
tylus wirshingi, Gymnodactylus geckoides, 
Thecadactylus rapicauda, and Tarentola maur- 
itanica (Phyllodactylidae). Our sampling out- 
side sphaerodactylids is minimal considering 
the diversity of the Gekkota as a whole and 
we recognize that taxon sampling may 
significantly affect tree topology. However, 
as our focus is on the characters that define 
Sphaerodactylidae, sphaerodactyls, and Spha- 
erodactylus we believe that this limitation will 
not materially affect our results. Twenty 
independent searches were done using de- 
faults of "xmult" plus 10 cycles of tree drifting 
(Goloboff, 1999). Support was estimated 


No. 529 

Figure 2. Right lateral view of Sphaerodactylus ciguapa sp. nov. Scale bar = 5 cm. 

through Bremer support indices (BS; Bremer, 
1994), relative Bremer support indices (rela- 
tive fit difference [RFD]; Goloboff and Farris, 
2001), bootstraps, and symmetric resampling 
(SR) expressed as GC values (difference in fre- 
quencies for groups supported-contradicted; 
Goloboff etdL, 2003b). 

External and osteological features of addi- 
tional taxa of Sphaerodactylus and other 
sphaerodactylids (Appendix 2) were also 
compared in the course of diagnosing the 
new species, but were not incorporated into 
the phylogenetic analysis. Original descrip- 
tions and other references (Schwartz and 
Graham, 1980; Schwartz and Thomas, 1983; 
Schwartz and Henderson, 1991) were consult- 
ed for further information about body size 
and scalation features of Sphaerodactylus spp. 

Institutional Abbreviations. Material exam- 
ined was obtained from the following collec- 
tions: Aaron M. Bauer personal collection, 
Villanova University, Villanova; (AMB); 
American Museum of Natural History, New 
York (AMNH); The Natural History Muse- 
um, London (BMNH); California Academy 
of Sciences, San Francisco (CAS); Field 

Museum of Natural History, Chicago 
(FMNH); James Ford Bell Museum, Univer- 
sity of Minnesota, - Saint Paul (JFBM); 
Museum of Comparative Zoology, Harvard 
University, Cambridge (MCZ); Museum of 
Vertebrate Zoology, University of California, 
Berkeley (MVZ), Museu de Zoologia, Uni- 
versidade de Sao Paulo, Sao Paulo (MZUSP); 
Sam Noble Oklahoma Museum of Natural 
History, University of Oklahoma, Norman 
(OMNH); Richard Thomas personal collec- 
tion, Universidad de Puerto Rico, Rio Pie- 
dras, Puerto Rico (RT); Museo de Zoologia, 
Universidad de Puerto Rico, Rio Piedras 
(UPRRP); United States National Museum, 
Washington (USNM; USNMFH— Field Se- 
ries); Coleccion de Herpetologia de la Uni- 
versidad del Valle, Cali, Colombia (UV-C). 
Description of new species 
Sphaerodactylus ciguapa Daza and Bauer, 

new species 

Figures 2-7 

Holotype. MCZ R- 186380, amber-embedded, 
nearly complete skeleton with patches of 
integument. Collected from the late Early 
Miocene to early Middle Miocene (15 to 




pelvic girdle ri 9 ht P es 

right humerus 


pelvic girdle 


— jaw 

— scapulocoracoid 

left humerus 

pelvic girdle 

left pes 

left radius 

left ulna 

left manus 

right humerus 


left arm 

Figure 3. (A) Dorsal, (B) lateral, and (C) ventral X-rays of Sphaerodactylus ciguapa sp. nov. Scale bar = 5 cm. 
The partially radiopaque "V"-shaped element in the caudal area labeled with a question mark may represent a 
portion of the regenerated tail or an artifact unrelated to the specimen itself. 

20 MYA) amber deposits of La Toca mine, 
in the Cordillera Septentrional, Santiago 
Province, north of Municipio Santiago de 
los Caballeros, Dominican Republic. - 

Diagnosis. A medium-sized Sphaerodacty- 
lus with an estimated snout-vent length 
(SVL) of 33 mm. Basicranium with narrow 

clinoid process; rounded crista alaris; 
straight crista prootica; squarish paroccipital 
process; knoblike sphenoccipital tubercle; 
fenestra ovalis completely visible in ventral 
view; foramen magnum roughly oval. Clav- 
icles each with a single enlarged fenestra; 
interclavicle with broad lateral arms: 26 


No. 529 

Figure 4. Dorsal view of basicranium, jaw, hyoid apparatus, and atlas of Sphaerodactylus ciguapa sp. nov. Gray 
areas with white zig-zags indicate portions of the specimen worn during polishing. Abbreviations: lcb, first 
ceratobranchial; 2cb, second ceratobranchial; at, atlas; bhy, basihyal; bo, basioccipital; bp, basipterygoid process; 
cal, crista alaris; clp, clinoid process; cob, compound bone (angular, articular, prearticular); cor, coronoid; crs, crista 
sellae; ept, epipterygoid; fco, fossa columellae, hhy, hypohyal; mf, mandibular fossa; occ, occipital condyle; oto, 
otooccipital; pop, paroccipital process; ppp, postparietal process; pro, prootic; pt, pterygoid; q, quadrate; sa, 
surangular; set, sella turcica; pbsph, parabasisphenoid; sq, squamosal; tbr, trabeculae. Scale bar = 5 mm. 

presacral vertebrae; pelvis with large and 
ventrally directed pectineal process; digits 
short, with manual metacarpals twice the 
length of the phalanges; fourth phalangeal 
element of the fourth manual digit short. 
Gular and body laterodorsal scales small, 
rounded posteriorly, and juxtaposed to weak- 
ly imbricate; some lateral scales distinctly 
keeled; forelimb scales smooth and strongly 
imbricate; claw enclosed by three scales. 

Ninety-nine extant and one fossil species 
of Sphaerodactylus are currently recognized 
as valid (Bohme, 1984; Kluge, 2001; Uetz, 
2011). In general, Sphaerodactylus are 
known as endemics of small areas (Schwartz 
and Henderson, 1991; Henderson and 
Powell, 2009); because of this it is reason- 

able to compare this fossil with the 35 
extant species from Hispaniola, as well as 
the other fossil species. However, because 
the age of the Dominican amber deposits 
is older than, or contemporaneous with, 
estimations of the formation of the Mona 
Passage and the separation of Hispaniola 
and Puerto Rico (-16-11 MYA; Iturralde- 
Vinent and MacPhee, 1996; MacPhee et al., 
2003) we also compared the. new species 
with 10 extant species from the "Puerto 
Rico Area" (sensu Thomas, 1999), an area 
that includes the islands of Mona, Monito, 
and Desecheo as well as Greater Puerto 
Rico (sensu Thomas and Schwartz, 1966) 
(Appendix 2; species endemic to St. Croix, 
U.S. Virgin Islands were not included as 




Figure 5. Right lateral view of Sphaerodactylus ciguapa sp. nov. showing skull, cervical vertebrae, and pectoral 
girdle. Gray areas with white zig-zags indicate portions of the specimen worn during polishing. Abbreviations: lcb, 
first ceratobranchial; at, atlas; ax, axis; civ, clavicle; cob, compound bone; cor, coronoid; cv#, cervical vertebrae #; 
epi, epipterygoid; h, humerus; hvc, groove for the course of the lateral head vein; hy, hypapophyses; mf, mandibular 
fossa; ocr, occipital recess; ppp, postparietal process; pt, pterygoid; rap, retroarticular process; rib, rib; sco, 
scapulocoracoid; scofo, scapulocoracoid foramen; sq, squamosal; V, incisura prootica for the course of the trigeminal 
nerve. Scale bar = 5 mm. 

this island is not part of the Puerto Rican 

The specimen of S. ciguapa is skeletally 
mature (see Discussion), and is comparable 
in size (here we have considered 29-36-mm 
SVL to be in the same size range of S. 
ciguapa) to 1 7 extant species from Hispaniola 
(S. altavelensis, S. armstrongi, S. asterulus, S. 
cinereus, S. clenchi, S. darlingtoni, S. diffici- 
lis, S. lazelli, S. leucaster, S. randi, S. 
rhabdotus, S. samanensis, S. savagei, S. 
schuberti, S. shrevei, S. thompsoni, and 
S. zygaena), four from the Puerto Rico Area 
(S. monensis, S. klauberi, S. macrolepis, S. 
micropithecus), and to the amber-preserved 
species S. dommeli (Bohme, 1984). Of these 
22 species, S. ciguapa may be distinguished 
from S. monensis, S. macrolepis, and S. 
thompsoni by its much smaller dorsal scales, 
from S. samanensis by its larger and more 
swollen scales, from S. cinereus by its 
heterogeneous dorsal scalation including 
imbricating, keeled scales (versus granular 
dorsal scalation), and from all others except 

S. asterulus, S. difficilis, S. dommeli, S. 
rhabdotus, and 5. shrevei by its swollen, 
weakly keeled to keelless dorsal scales 
(versus flat scales with strongly to very 
strongly keeled scales, see Fig. 8 for exam- 
ples of scale features discussed). The new 
species may be distinguished from S. rhab- 
dotus by its more weakly keeled and 
subimbricate (versus strongly keeled and 
strongly imbricate) dorsal scales, and from 
S. asterulus, S. difficilis, and S. shrevei by the 
presence of an extremely large clavicular 
fenestra (versus a small fenestra). The amber- 
embedded S. dommeli, which comes from the 
same mine area as S. ciguapa, may be 
differentiated on the basis of its smaller, 
more granular dorsal scales (see Bohme, 
1984, fig. 3). Additionally, we were unable 
to identify enlarged clavicular fenestrae in 
X-rays of S. dommeli. 

General Description. The fossil is enclosed 
in an oval piece of polished amber measuring 
48.5 mm in its maximum dimension. The 
specimen lies close to one of the margins of 


No. 529 

Figure 6. Sphaerodactylus ciguapa sp. nov. in dorsolateral view showing portions of the pectoral girdle. Scale 

bar = 2 mm. 

the piece and in some spots it is exposed as a 
result of the polishing process. The amber 
has a partial fracture plane at the posterior 
end of the specimen, but the two portions 
remain together. The amber matrix embed- 

ding the specimen is semitransparent yellow 
and the bone color is dark brown, providing 
a contrast that facilitates observation oi" the 
whole specimen (Fig. 2). The specimen is 
mainly skeletonized (Figs. 2, 3), with a few 



Figure 7. Sphaerodactylus ciguapa sp. nov. showing (A) left laterodorsal scapular integument, (B) mid-trunk 
dorsal scales adjacent to the vertebral line, (C) left hand showing the typical Sphaerodactylus scalation pattern 
around the claw, and (D) dorsal scales of S. difficilis (USNM 328965), an extant species from Hispaniola with similar 
scalation to S. ciguapa. Scale bar = 0.5 mm. 

scattered patches of skin. The skeleton 
preserves the posterior half of the left 
pterygoid, a portion of the left epipterygoid, 
a tiny fragment of the left parietal, left 
squamosal, left quadrate, and some portions 
of the brain case (including left prootic, left 
otooccipital, parabasisphenoid, and basioc- 
cipital), the posterior part of the right 
mandibular ramus, parts of the hyoid appa- 
ratus, all of the cervical, thoracolumbar, 
sacral, pygal, and some postpygal caudal 
vertebrae, the left arm and the proximal 
portion of the right humerus, both suprasca- 
pulas, scapulocoracoids, and clavicles, the 

complete sternum, the pelvis, left femur and 
tibia, and both feet. All elements, except the 
right pes, are articulated. 

Hob type Measurements (unless otherwise 
stated, measurements were made along the 
long axis of each element; for paired 
elements, left side measurement is provided): 
braincase from the tip of the basipterygoid 
process to the occipital condyle: 3.66 mm; 
quadrate length from the cephalic condyle to 
the mandibular condyles: 1.9 mm; squamosal: 
0.71 mm; jaw fragment: 3.28 mm; cervical + 
thoracolumbar vertebrae: 24.91 mm; sacrum 
length: 0.88 mm; sacrum width: 2.31 mm; tail 



No. 529 

Figure 8. Dorsal scale variation in Sphaerodactylus geckos. (A) S. pacificus (USNM 157531, small, rounded, 
swollen, juxtaposed), (B) S. rosaurae (USNM 570205, medium, rounded, swollen, juxtaposed, with a middorsal zone 
of granular scales), (C) S. parkeri (USNM 328281, large, rounded, not swollen, imbricate), (D) S. argus (USNM 
251978, small, acute, not swollen, imbricated), (E) S. richardsoni (USNM 252126, large, acute, swollen, imbricate), 
(F) S. thompsoni (USNM 328977, large, rounded, swollen, juxtaposed). 

(proximal segment only preserved): 3.77 mm; 
scapulocoracoid from the fossa glenoidea 
to the dorsal margin: 2.28 mm; humerus: 
4.46 mm; ulna: 3.23 mm; radius: 2.79 mm; 
third manual digit + metacarpal: 2.43 mm; 
pelvic girdle from the epipubic cartilage to the 
posterior edge of illium: 3.31 mm; metischial 
process: 0.35 mm; tibia: 2.77 mm; third pedal 
digit + metatarsal: 3.31 mm. 

Dermatocranium. The parietal is only 
represented by the tip of the posterior end 
of the left postparietal process (ppp, Figs. 4, 
5); the postparietal process is very narrow 
and contacts the squamosal laterally at the 

midpoint of this bone, as in other sphaero- 
dactyls. The squamosal (sq, Figs. 4, 5) is 
small, slightly curved, and rounded in cross- 
section. Its distal end contacts the braincase 
and the top of the quadrate. A fragment of 
the left pterygoid (pt, Figs. 4, 5) extends 
from a point anterior to the fossa columellae 
(fco, Fig. 4) to the end of the quadrate pro- 
cess. Basispterygoid-pterygoid, epipterygoid- 
pterygoid, quadrate-pterygoid, and cranio- 
mandibular skull joints are preserved. The 
former two are synovial and the latter a 
syndesmosis (Frazzetta, 1962; Payne et al, 
201 1). In lateral view the pterygoid is mostly 




straight; it possesses a large facet for the basis- 
pterygoid joint that is visible in medial view. 

Splanchno cranium. A nearly complete left 
epipterygoid (ept, Figs. 4, 5) is preserved, 
although the dorsal portion of this bone is 
broken and is disarticulated from the pro- 
otic. Only the left quadrate bone (q, Figs. 3, 
4) is preserved. The bone is completely 
convex and the dorsal margin is rounded 
with no lateral indentation. Although the 
craniomandibular joint is in situ, it can be 
seen that the distal articular surface bears 
two condyles, and that the lateral is slightly 
larger than the medial one. The presence and 
position of the quadrate foramen cannot be 
established in the specimen. The left auditory 
meatus is nearly intact, and includes portions 
of the tympanic membrane, implying that the 
left stapes is preserved within the middle ear, 
although it is not visible. 

Neurocranium. The parabasisphenoid com- 
plex (pbsph, Fig. 4) is fused posteriorly 
to the basiocciptal. The left basipterygoid 
process (bp, Fig. 4) is short but expanded 
distally and anterolateral^ oriented. It is 
partially covered by a long, narrow clinoid 
process (dp, Fig. 4) that roofs the notch on 
the basipterygoid process and marks the 
course of the lateral head vein (hvc, Fig. 5). 
The right basipterygoid process is missing. 
The paired trabeculae (tbr, Fig. 4) are 
clearly distinguishable; they are round in 
cross-section and parallel to one another. 
Posterior to the trabeculae, the sella turcica 
(set, Fig. 4) is bounded posteriorly by an 
anteriorly curved crista sellae (crs, Fig. 4). 
The anterior opening of the Vidian canal is 
located ventral to the crista sellae. Most of 
the basioccipital (bo, Fig. 4) is preserved; it 
is concave dorsally and forms part of the 
double occipital condyle (occ, Fig. 4) and 
the ventral border of the foramen magnum. 
The sphenoccipital tubercle epiphysis is 
small and knoblike and is located anteriorly, 
causing the crista tuberalis to be inclined 

posterodorsally. The left prootic (pro, 
Fig. 4) is preserved but its medial surface 
is partly worn down because of the polishing 
process. The crista alaris is rounded and 
small, and does not overhang the inferior 
process of the prootic. The inferior process 
bears the incisura prootica, which in geckos 
is closed, forming an oval foramen that 
surrounds a portion the mandibular branch 
of the trigeminal nerve, CN5 (V, Fig. 5). A 
portion of the left otooccipital (oto, Fig. 4) 
is present, but none of the foramina in the 
occiput (i.e., vagus and hypoglossal foram- 
ina) are discernable. 

Mandible (Figs. 3-5). The posterior por- 
tion of the left jaw comprises the posterior 
process of the coronoid (cor), the surangular 
(sa), compound bone (cob; angular, prear- 
ticular and articular), and, on the labial side, 
the posterior portion of the dentary. A wide 
mandibular fossa is formed by the suran- 
gular and the compound bone. This fossa 
opens laterally through a small slit (external 
mandibular fenestrae) that marks the sepa- 
ration between the partially fused surangu- 
lar and the compound bone (Daza et al., 

Hyoid Apparatus (Figs. 4, 5). A portion of 
the basihyal (without the glossohyal process) 
is preserved. The second epibranchials are 
articulated to the basihyal and oriented 
almost parallel to one another, as in S. 
macrolepis (Noble, 1921). Both first epibran- 
chials are preserved and curve upward 
toward the posterior portion of the brain- 
case. Anterior to the right second cerato- 
branchial there is an elongated bony struc- 
ture that could be a portion of the right 

Vertebral Column. All the presacral verte- 
brae are preserved. The total number is 26, 
as is typical for most geckos. There are eight 
cervical vertebrae (Fig. 5), which follows the 
commonest formula for lizards: 3 (ribless) + 
3 (short distal widened ribs) + 2 (long slender 



No. 529 

ribs) = 8 (Hoffstetter and Gasc, 1969). The 
intercentra of the atlas, axis, and third-sixth 
cervicals bear ventral hypapophyses and are 
positioned intervertebrally, remaining un- 
fused from the vertebrae centra (type A, 
Hoffstetter and Gasc, 1969). The hypapo- 
physes are double in the atlas and axis and 
single in the remaining cervicals. The orien- 
tation of the hypapophyses varies ventrally 
(atlas), posteriorly (axis), and anteriorly 
(remaining cervicals). The height of the 
seventh and eighth cervicals is 25% greater 
than that of the anterior cervicals, having 
taller and squarer neural arches when viewed 
laterally. These last two cervical vertebrae 
are more similar to the thoracolumbar series. 
All vertebrae bearing ribs have synapophyses 
(parapophysis + diapophysis; Hoffstetter and 
Gasc, 1969) that project laterally from the 
anteroventral part of the centrum (Fig. 5). 
The short ribs of cervical vertebrae 4-6 are 
not bifurcated distally (cv4-cv5, Fig. 5), but 
in sphaerodactyls, these ribs have a cartilagi- 
nous terminus that is not preserved in the 
fossil. Ribs from the fourth and fifth 
cervicals are free, and the sixth and seventh 
contact the medial surface of the scapulocor- 
acoid. The dorsal process of the clavicle 
contacts the dorsal surface of the sixth 
vertebral rib. Four or five vertebrae are 
connected to the sternum via sternal ribs 
(Fig. 3). The remaining thoracic vertebrae 
have long ribs that decrease in length 
posteriorly, each bearing small postxiphi- 
sternal inscriptional ribs. There is one ribless 
lumbar vertebra. The sacrum has fused 
transverse processes; the first sacral has an 
expanded transverse process that overlaps 
the second sacral. The exact number of pygal 
vertebrae (i.e., caudals lacking chevrons; 
Russell, 1967; Hoffstetter and Gasc, 1969) 
could not be determined, but there are at 
least five caudal vertebrae with elongated 
transverse processes. The tail seems to be 
regenerated, appearing as a poorly defined 

cartilaginous rod that is broken and bent and 
is situated along the posterior portion of the 
body (?, Fig. 3). 

Pectoral Girdle and Forelimbs (Fig. 6). The 
two clavicles are expanded medially, rotated 
forward, and articulated medially, contact- 
ing the anteroventral end of the interclavicle. 
The clavicles each have a single fenestra, 
which is among the largest seen in any 
sphaerodactyl examined. 

Pelvic Girdle and Hind Limbs (Fig. 3). The 
pelvic girdle and hind limbs are mainly 
covered by integument and are only visible 
in the X-rays. The ischium, pubis, and ilium — 
which is articulated with the sacrum — are 
fused. The two inominate bones are still 
articulated at the pubic (epipubic cartilage 
preserved) and ischial symphyses, forming a 
large ischiopubic fenestra (Figs. 3A, C). The 
pectineal process of the pubis is large and 
ventrally directed, as in all sphaerodactyls 
(Noble, 1921; Gamble et at., 2011a). The 
posterior flange of the ischium is more or less 
straight. The left acetabulofemoral joint is 
preserved; the left leg retains all of its 
elements. Of the right hind limb, only the 
pes, which is twisted and facing the front of 
the pelvis, is preserved. An exact phalangeal 
formula cannot be determined from the X- 
rays because of the superposition of the 
vertebrae, but all Sphaerodactylus known have 
manual and pedal formulae of 2:3:4:5:3 
and 2:3:4:5:4, respectively, with phalanges 2 
and 3 of digit 4 of both manus and pes 
shortened (Russell and Bauer, 2008). 

Integument. There are scales present in the 
gular, dorsolateral trunk, and apendicular 
regions. Gular scales are small, flattened, 
rounded posteriorly, not swollen, and juxta- 
posed; lateral scales covering the scapular 
blade and the body flanks are small, moder- 
ately keeled, rounded to subacute posterior- 
ly, slightly swollen, and juxtaposed with little 
or no imbrication (Fig. 7A); dorsal scales on 
the mid-trunk are slighty larger, unkeeled to 




weakly keeled, oval, slightly swollen, and 
subimbricate to weakly imbricate (Fig. 7B). 
The scales covering the forelimbs are round- 
ed posteriorly, smooth and strongly imbri- 
cated; the claw is enclosed by three scales, 
which are arranged in the typical asymmet- 
rical pattern of Sphaerodactylus (Fig. 7C): an 
enlarged outer inferolateral, a terminal + 
median dorsal, and an inner inferolateral 
{sensu Parker, 1926), or ventral, dorsal, and 
ventrolateral (sensu Kluge, 1995). 

Etymology. "La Ciguapa" is a Spanish 
name for a mythical humanoid of Domini- 
can folklore. It is described as a woman with 
brown or dark blue skin, whose feet face 
backward, and who has a very long mane of 
smooth, glossy hair that covers her naked 
body (Angulo Guridi, 1866; Perez, 1972; 
Ubinas Renville, 2000, 2003). It is supposed 
to inhabit the high mountains of the 
Dominican Republic. The name, treated here 
as a noun in apposition, recalls the dark 
brown bones and twisted feet in the holotype 
specimen and the source of the specimen in 
the Cordillera Septentrional of the Domini- 
can Republic. The Ciguapa legend has been 
proposed to be derived from the "opias" 
(spirits of the dead) of the indigenous 
Caribbean Taino people (Bosch Gavifio, 


Phylogenetic Analysis. Rooting with any 
of the outgroup taxa resulted in the same 
ingroup topology and measures of support, 
so Hemidactylus brookii was arbitrarily 
chosen to root all trees presented herein. 
Tree searches found four most parsimonious 
trees (MPT) of 1,135 steps (Consistency 
index [Ci] = 0.379. Retention index [Ri] = 
0.404). Two of these trees recovered a 
monophyletic Sphaerodactylidae, as strongly 
supported by molecular data (Gamble et al. , 
2008a, 2008b, 2011b); therefore we used one 

of these trees as working hypothesis for 
mapping characters. Nine internal nodes are 
well supported, with absolute BS values 
greater than or equal to 3, and relative BS 
values above 1 1 . Five of these nine nodes 
also had bootstrap values above 82 and GC 
values above 77 (Fig. 9). 

In the selected topology Aristelliger lar + 
Teratoscincus scincus are sister to remaining 
sphaerodactylids, with Quedenfeldtia, Eu- 
leptes, and Saurodactylus as sequential sister 
taxa to the clade formed by sphaerodactyls + 
Pristurus. Among the most parsimonious 
trees the positions of Lepidoblepharis and 
Sphaerodactylus are interchangeable within 
the sphaerodactyls. 

Character Mapping. As previously noted, 
there are only two nonmolecular characters 
that currently serve to diagnose the family 
Sphaerodactylidae, and neither of these is 
expressed in all members of the group or is 
exclusive to the clade. Thus, the monophyly 
of this family has not yet been tested using 
morphological data. Here we present all the 
characters that apply to each of three nested 
clades on the basis of their mapping on the 
preferred most parsimonious tree. For each 
one of named clades we emphasize those 
characters that exhibit less homoplasy and 
may therefore be useful for the morpholog- 
ical diagnosis of these groups. Characters 
that could be scored on S. ciguapa are 
indicated in bold numbers. 

Sphaerodactylidae: This clade inludes all 
the genera listed in Gamble et al (2008a) and 
is supported by seven characters: 10) convex 
snout; 136) dagger-shaped anterolateral pro- 
cess of frontal; 308) anterior inferior alveolar 
foramen surrounded by dentary, splenial, 
and angular; 334) anterior tip of splenial 
narrow and pointed; 439) branched xiphi- 
sternum; 506) metatarsal V greatly hooked 
(see Discussion); 560) two pygial vertebrae. 
Of these characters, 506 was not present in 
any other sampled gekkotan, whereas char- 



No. 529 

Hemidactylus brookii 
Phyllodactylus wirshingi 
Gymnodactylus geckoides 
Thecadactylus rapicauda 
Tarentola mauritanica 
Narudasia f estiva 
Pseudogekko smaragdinus 
Aristelliger lar 
Teratoscincus scincus 
Quedenfeldtia trachyblepharus 
Euleptes europaea 
Saurodactylus mauritanicus 
Pristurus carteri 
Gonatodes albogularis 
Lepidoblepharis xanthostigma 
Sphaerodactylus roosevelti 
Sphaerodactylus klauberi 
Sphaerodactylus ciguapa 
Coleodactylus brachystoma 
Chatogekko amazonicus 
Pseudogonatodes guinanensis 

Figure 9. One of four most parsimonious trees of gekkonoid geckos. Circles at nodes denote BS/RFD equal to 
or higher than 3/11, filled circles further indicate boostrap/GC values higher than 82/78. X-rays of representative 
sphaerodactylid geckos from the tree are shown at right (not to same scale). Initials next to each X-ray correspond to 
the the genus and specific epithet of taxa represented on the tree. 

acters 136 and 334, although present in all 
sphaerodactylids, are also found in some 

Sphaerodactylini + Pristurus: This clade is 
supported by 11 characters: 19) fenestra 
vomeronasalis continuous within the fenes- 
tra exochoanalis; 37) ascending nasal process 
of the premaxilla separates nasals through- 
out approximately half their length; 93) 
postorbitofrontal large, with no reduction 
of processes; 97) postorbitofrontal ventrolat- 
erally curved; 140) brief contact between the 
frontal and the maxilla; 266) fusion of 
parabasisphenoid and basioccipital; 332) 

splenial fused to the coronoid; 352) suran- 
gular contacts dentary posterior to the 
coronoid-dentary suture; 440) mesosternal 
extension absent; 454) humeral ectepicondyle 
continuous, consolidated with the bone 
shaft; 601) nostril in contact with rostral 
scale. The least homoplasic character was 
601, which is present only in Aristelliger 
outside sphaerodactyls. Although there were 
no exclusive characters for this clade, char- 
acters 93, 352, 440, and 454 were also 
invariably present among sphaerodactyls, 
but these character states occur in other 
sampled genera. 




Sphaerodactyls: This clade is equivalent 
to Sphaerodactylini (Gamble et al, 2008a). 
Althought this clade was not recovered in 
any of our MPTs, we performed a con- 
strained search forcing this New World 
clade, which receives strong molecular sup- 
port, to be monophyletic. We obtained a 
single MPT three steps longer than the 
shortest trees from the unconstrained analy- 
sis (1,138 steps; Ci = 0.353; Ri = 0.333). 
Eleven characters support this clade in the 
constrained analysis: 19) fenestra vomerona- 
salis continuous within the fenestra exochoa- 
nalis; 110) lacrimal foramen bounded by 
prefrontal and maxilla; 165) parietal nuchal 
fossa present and extending substantially 
onto the skull table; 184) posterior end of 
squamosal not in contact with dorsum of 
quadrate; 201) secondary palate formed 
around choanal groove of palatine, ventro- 
medial fold partly hides or hides most of the 
the choanal groove; 220) anterior point of 
the ectopterygoid relatively wide, abruptly 
tapering to point; 281) crista prootica of 
prootic with straight margin (except in 
Chatogekko, which has a triangular crista 
prootica); 340) coronoid low, hardly elevated 
above jaw outline; 486) pectineal process of 
pubis large and ventrally directed; 601) nostril 
in contact with rostral scale; 612) supraciliary 
spine present. Of these characters 201 and 340 
were not present in any other sampled 
gekkotan. Other characters that show low 
homoplasy are 165 and 601 (also in Aristelli- 
ger), 184 (also in Teratoscincus), and 486 (also 
in Thecadactylus). Character 612 is also 
present in Aristelliger but lost in Chatogekko, 
Coleodactylus, and Pseudogonatodes. 

Sphaerodactylus: Sixteen characters sup- 
port this genus: 8) anterorbital portion of the 
skull equals 30% or less of the total skull 
length; 10) flat snout; 19) fenestra vomefona- 
salis and incisura jacobsoni separated; 28) 
foramen magnum roughly oval; 98) postorbi- 
tofrontal, with large lateral process; 179) 

postparietal process length less than half the 
length anterior to the parietal notch; 330) 
presence of angular and surangular processes 
of dentary; 347) the anterolingual process of 
the coronoid separates the dentary and 
splenial anteriorly; 368) 12-13 premaxillary 
teeth; 586) body with keeled scales; 600) two 
to four loreal scales; 605) tympanic edge not 
smooth; 638) digits with the distal-most 
superolateral scales in contact; 651) dorsal 
color pattern of head and nape with light 
stripes; 652) dorsal color pattern of body with 
ocelli; 671) ear partially occluded by flaps of 
skin. Of the characters listed, 330, 638, 651 
were not found in any other sampled gekko- 
tan. Sphaerodactylus geckos differ from other 
sphaerodactyls by characters 19 and 605. 
Other characters that were less homoplastic 
were 179 (also present in Teratoscincus and 
Chatogekko), 368 (also present in Pseudogo- 
natodes), 586 and 600 (also present in Chato- 
gekko), and 671 (also present in Pristurus). 


Phytogeny. Four main hypotheses exist for 
the relationships of sphaerodactylid geckos 
(Fig. 10), two of them morphologically de- 
rived (Kluge, 1995; Arnold, 2009) and the 
others based on multigene analyses (Gamble 
et al, 2008a, 2011b). There are discrepancies 
in the branching pattern between the mor- 
phological and molecular topologies, mainly 
with respect to the basal relationships within 
Sphaerodactylidae. Whereas the molecular 
phylogenies (Gamble et al, 2008a, 2011b; 
Figs. 10C, D) include a clade comprising 
Pristurus, Euleptes, Teratoscinus, Aristelliger, 
and Quedenfeldtia, in the morphological hy- 
potheses Pristurus was found to be either the 
sister taxon of all sphaerodactyls (Kluge, 1995; 
Daza, 2008) or Pristurus + Quedenfeldtia 
were sister to sphaerodactyls (Arnold, 2009). 
The sister group relationship between Aris- 
telliger and Quedenfeldtia as suggested by 



No. 529 










































Figure 10. Phylogenetic relationships of sphaerodactylid geckos on the basis of previously published analyses. 
(A) Reanalysis of Kluge's (1995) morphological data set, (B) Arnold (2009), (C) Gamble et til. (2008a), and (D) 
Gamble et dl. (201 lb). "Sphaerodactyls" in Figure 10B includes all the miniaturized New World sphaerodactylids (i.e.. 
Coleodactylus, Gonatodes, Lepidoblepharis, Pseudogonatodes, Sphaerodactylus, and the newly recognized Chatogekko). 

multigene phylogenies is not congruent with 
our morphological results, in which an 
Aristelliger + Teratoscinus clade is supported 
by 14 morphological characters, one of them 
being the parafrontal bones, which are unique 
osteological structures in the circumorbital 
series only known in these two genera (Bauer 
and Russell, 1989; Daza and Bauer, 2010). 
The two molecular hypotheses differ mainly 
in the degree of resolution outside Sphaer- 
odactylinae and in the placement of the 
extremely modified Coleodactylus amazonicus 
group (Figs. 10C-D), which has recently been 
recognized as a new sphaerodactyl genus, 
Chatogekko (Gamble et a/., 201 la). 

The branching pattern we obtained for the 
sphaerodactyl clade is consistent with a 

previous morphological hypothesis (Kluge, 
1995; Fig. 10A), but there is also a degree of 
taxonomic congruence between our hypothe- 
sis and recent multigene phylogenies (Gamble 
et al, 2008a, 2011b). For instance, both 
molecular and morphological data provide 
strong support for the Sphaerodactylinae (i.e., 
sphaerodactyls + Saurodactylus), although 
they differ in the position of Pristurus. 

Previous morphological analyses have not 
indentified synapomorphies that support 
Sphaerodactylidae; for instance, a reanalysis 
of Kluge's (1995) data set using the gekkonid 
genus Cnemaspis to root the tree results in a 
MPT in which Narudasia (another gekkonid) 
is nested within Sphaerodactylidae (Fig. 10A). 
Our new analysis including a superior number 




of characters (approximately 27 and 55 times 
the number of morphological characters of 
Kluge [1995] and Arnold [2009], respectively) 
provides provisional empirical morphological 
evidence for the monophyly of Sphaerodacty- 
lidae and two clades nested within it. Although 
characters or combinations of characters 
support the monophyly of less inclusive clades 
like Sphaerodactylus and sphaerodactyls, rela- 
tively homoplasy-free characters supporting 
the Sphaerodactylidae remain elusive. The 
reduction of clutch size from two to one (see 
Gamble et al, 2008a) remains a possible 
synapomorphy for the family, although on 
the basis of our topology (Fig. 9) it is equivocal 
if this character applies at the level of the 
Sphaerodactylidae as a whole, or to this clade 
exclusive of Teratoscincus + Aristelliger. 

A strongly hooked metatarsal V was 
the least homoplastic trait supporting the 
Sphaerodactylidae in our analysis. Although 
this was not seen in any of the outgroup taxa 
in our phylogenetic analysis, this character is 
not exclusive to the Sphaerodactylidae, as 
both straight (Figs. 11A-C) and hooked 
(e.g., Ailuronyx seychellensis, Fig. 11D) mor- 
phologies occur in other gekkonoids. Among 
the Sphaerodactylidae this bone is variable 
but is always bent, being strongly hooked in 
some genera (e.g., Aristelliger, Quedenfeldtia, 
Teratoscincus; Figs. 11E-G) or more gently 
curved (e.g., sphaerodactyls, Fig. 11H). 

Despite the fragmentary nature of S. 
ciguapa it was possible to score it for 258 
characters (38.2% of the complete list). The 
analysis of these data unambiguously sup- 
ports its placement within the genus Sphaer- 
odactylus. Unfortunately in our phylogenetic 
analysis Sphaerodactylus was represented by 
only a few species from the argus series from 
Puerto Rico; hence this hypothesis is not 
useful for establishing the intrageneric rela- 
tionships of S. ciguapa. Our comparisons 
with living taxa from Hispaniola and Greater 
Puerto Rico (see Diagnosis) suggest at least 

phenetic similarity with S. difficilis, a mem- 
ber of a widespread and diverse species 
complex (Thomas and Schwartz, 1983) in 
the notatus species group of the argus series, 
and with members (S. shrevei, S. asterulus, 
S. rhabdotus) of the shrevei species group 
(Schwartz and Graham, 1980) in the cinereus 
series. However, as these two groups span 
most of the phylogenetic diversity within 
West Indian Sphaerodactylus (Hass, 1991, 
1996), the more specific affinities of S. 
ciguapa remain uncertain. 

Morphology. The skeletal anatomy of at 
least some representative Sphaerodactylus 
geckos has been studied in detail (Noble, 
1921; Parker, 1926; Daza et al, 2008). In 
conjunction with the new data derived from 
S. ciguapa it is possible to reevaluate certain 
aspects of the osteology of the genus, and 
sphaerodactyls more broadly, within the 
more inclusive framework of the Gekkota. 

The clinoid process of the parabasisphe- 
noid is variable. In this fossil it is narrower 
than that described in S. roosevelti (Daza 
et al, 2008), or observed in S. difficilis; it is 
unknown how variable this structure is 
across Sphaerodactylus species, but it might 
be a diagnostic character at some level. The 
paired (unfused) trabeculae in Sphaerodacty- 
lus are connected by a bony lamina in adults, 
including the type of S. ciguapa, whereas in 
juveniles these are discrete (Daza et al., 
2008). The sphenoccipital tubercle is reduced 
in small sphaerodactyls (excluding Gona- 
todes) and a similar reduction is present in 
miniaturized lizards from all gekkotan fam- 
ilies (e.g., Aprasia [Pygopodidae], Coleonyx 
[Eublepharidae], Narudasia [Gekkonidae], 
Homonota [Phyllodactylidae]). The reduction 
of the apophysis that caps the sphenoccipital 
tubercle suggests modifications to the tendi- 
nous attachment of the fourth division of the 
m. longissimus capitis, which extends back 
into the ventral neck region along the fourth 
cervical in lepidosaurs (Al Hassawi, 2007) 


No. 529 


Figure 1 1 . Left pes of some gekkonoid lizards showing variation on the shape of metatarsal V (shaded in gray) 
in phyllodactylids (A, B), gekkonids (C, D), and sphaerodactylids (E-I). (A) Phyllodactylus wirshingi (CAS 175498). 
(B) Tarentola mauritanica (UC MVZ 178184), (C) Pseudogekko brevipes (CAS 128978), (D) Ailuronyx seychellensis 
(CAS 8421, (E) Aristelliger lav (USNM 260003), (F) Quedenfeldtia trachyblepharus (USNM 196417). (G) 
Teratoscincus scincus (CAS 101437), (H) Lepidoblepharis xantostigma (USNM 313791), and (I) Sphaerodactylus 
klauberi (UPRRP 006416). 




and whose fibers are attached to the ventral 
hypapophyses of the cervical vertebrae. Since 
reduction of this tubercle seems to be present 
only in small species, it is possible this is 
a character linked to miniaturization and 
might be related to a reduction of the neck 
muscle fibers. 

In S. ciguapa, as in the rest of sphaer- 
odactyls, the basicranial elements are fused, 
obscuring the sutures between the braincase 
bones. Observations of juveniles and newly 
hatched Sphaerodactylus indicate that the 
fenestra ovalis is bounded anteriorly by the 
prootic and posteriorly by the otooccipital 
(Daza et al. , 2008); hence this fenestra serves 
to estimate the limit between these two 
elements. In gekkotans the otooccipital has 
a synchondrosis articulation with the quad- 
rate (Payne et al, 2011), although this joint 
also has been described as syndesmosis 
(Webb, 1951). In geckos the articulation of 
the quadrate has been described as "parocci- 
pital abutting" where this bone forms a well- 
defined articular process, which is applied 
against the anteroventral aspect of the 
paroccipital process (Rieppel, 1984). The 
paroccipital process of geckos has been 
described as thick or thin (Jollie, 1960), and 
this variation seems to be related to skull 
size, being generally elongated in larger 
species and reduced in small species. Similar 
variation has been seen in size series of 
amphisbaenians (Montero and Gans, 2008). 
Sphaerodactylus spp. have a small, thick 
paroccipital process that in posterior view 
is square (i.e., width and height subequal). 
Shape and size of the paroccipital process 
define its participation in the quadrate 
suspension in gekkotans. In Pristurus, Gona- 
todes, and Lepidoblepharis this process forms 
a true paroccipital abuttment, but in Sphaer- 
odactylus, Chatogekko, Coleodactylus, "and 
Pseudogonatodes, the quadrate is suspended 
from the lateral surface of the braincase, in 
front of the paroccipital process (pop, 

Fig. 4), with no participation of the squa- 
mosal. In very small forms (e.g., Chatogekko) 
the paroccipital process is so small that it has 
minimal or no participation in the suspen- 
sion of the quadrate (Gamble et al, 2011a). 

Ventral to the paroccipital process is 
located the rounded occipital recess, which 
in adult lizards represents the recessus scale 
tympani (Oelrich, 1956; Rieppel, 1985) and 
which in Sphaerodactylus is exclusively sur- 
rounded by the otooccipital (Daza et al, 
2008). In other gekkonomorphs participa- 
tion of the basioccipital in the margin of the 
occipital recess has been reported (Kluge, 
1962; Grismer, 1988; Conrad and Norell, 
2006). This participation is due to an out- 
growth of the sphenooccipital tubercle; there- 
fore the medial margin of the occipital recess 
is a good indication of the boundary between 
the otooccipital and the basiooccipital in 
forms with a fused braincase. 

The perforated stapes, commonly present 
among sphaerodactylids, is uncommon among 
squamates; this feature has only been reported 
in some gekkotans, some amphisbaenians, and 
dibamids (Greer, 1976; Kluge, 1983; Rieppel, 
1984; Gauthier et al, 1988, Bauer, 1990; 
Conrad, 2008; McDowell, 2008). The only 
sphaerodactylid where the stapes is unperfo- 
rated is Saurodactylus, where this bone is short 
and has a thick shaft and large footplate 
(Evans, 2008). Although in S. ciguapa this 
bone is not visible, it is very likely that it has a 
stapedial foramen like its congeners. 

The number of presacral vertebrae among 
sphaerodactylids is variable; most of the 
genera have the most common gekkotan 
number of 26 (Hoffstetter and Gasc, 1969), 
whereas in Quedenfeldtia and Pristurus this 
number is reduced to 24 and 23, respectively. 
Arnold (2009) related the reduction of verte- 
brae in Pristurus species to a shift from active 
foragers to ambush predators. He also scored a 
reduction of presacral vertebrae in Saurodac- 
tylus mauritanicus, which was not corroborat- 



No. 529 

Figure 12. Ventral view of the pectoral girdles of sphaerodactyl geckos. (A) Sphaerodactylus glaucus (MVZ 
Herps 149093), (B) Gonatodes albogularis (MVZ Herps 83369), (C) Lepidoblepharis xantostigma (USNM 313791), 
(D) Coleodactylus brachystoma (MZUSP uncatalogued), (D) Chatogekko amazonicus (AMNH R 132039), (E) 
Pseudogonatodes guinanensis (MZUSP 94826). Abbreviations: ace, anterior coracoid emargination; cf, clavicular 
fenestra; civ, clavicle; eco, epicoracoid; fg, fossa glenoidea; h, humerus; iclv, interclavicle; me, mesosternal extension; 
sco, scapulocoracoid; scof, scapulocoracoid fenestra; scofo, scapulocoracoid foramen; stn, sternum; stnr, sternal rib; 
xy, xiphisternum. Scale bar = 1 mm. 

ed by the specimens we reviewed. Reduction of 
presacral vertebrae explains the development 
of stocky bodies in Quedenfeltia and Pristurus, 
a process that according to the current 
morphological hypothesis and the molecular 
topologies would have been independently 
acquired (but see Arnold, 2009). 

Centrum morphology of presacral verte- 
brae is procoelic in most sphaerodactyls, 

whereas Gonatodes has amphicoelous verte- 
brae (Hoffstetter and Gasc, 1.969; Kluge, 
1967, 1995). The formation of procoelous 
vertebrae in Sphaerodactylus proceeds differ- 
ently from that in the rest of squamates. In 
these geckos the intervertebral tissue does 
not form a condyle but persists (Werner, 
1971), suggesting that the procoelous verte- 
brae in these geckos might be derived from 




amphicoelous ancestors (Hoffstetter and 
Gasc, 1969; Werner, 1971) with vertebrae 
resembling those of Gonatodes. Caudal ver- 
tebrae in sphaerodactyls have the first 
autotomy plane within the sixth or seventh 
caudal, as is consistent with S. ciguapa. It 
appears as if the type may have rebroken its 
tail at the proximal-most autotomy plane in 
an effort to escape when trapped in the resin. 

A clavicular fenestra seems to be a constant 
character for Sphaerodactylus (Noble, 1921; 
Gamble et al, 2011a), and this opening is 
huge in S. ciguapa. In Gonatodes, Lepidoble- 
pharis, and Chatogekko the clavicles are 
unperforated (Figs. 12B, C, E; Noble, 1921; 
Parker, 1926; Gamble et al, 2011a), where- 
as in Coleodactylus and Pseudo gonatodes 
these may be closed or open, and the pres- 
ence of a fenestra may be asymmetrical 
(e.g., Fig. 12D). The lateral arms of the 
interclavicle are another variable feature 
within sphaerodactyls (Kluge, 1995). In some 
Sphaerodactylus, including S. ciguapa, and in 
Chatogekko the arms are almost indistin- 
guishable, producing an almost rhomboid 
interclavicle (Figs. 6, 12A, E); in Gonatodes 
the interclavicle is cruciform with elongated 
arms (Fig. 12B; Rivero-Blanco, 1976, 1979); 
Lepidoblepharis have short, squarish arms 
(Fig. 12C); Coleodactylus has broad, rounded 
arms (Fig. 12D); and in Pseudogonatodes 
interclavicle shape is variable — P. barbouri 
has an interclavicle with rounded lateral arms 
(Noble, 1921; Parker, 1926), and in P. 
guinanensis the interclavicles have no lateral 
arms (Fig. 12F). Variation in the develop- 
ment of the lateral arms of the interclavicle is 
likely to be correlated with differences in the 
posterior insertion of the sternohyoid muscle. 

Although the appendicular skeleton pre- 
sents no obviously phylogenetically informa- 
tive characters, the epiphyses of the long 
bones are fused to the diaphyses, confirming 
that the type of S. ciguapa is skeletally 

Unfortunately scale descriptors have not 
been used consistently by different authors 
and variation, both between individuals and 
across the dorsum of single animals, can be 
extreme, rendering both the characterization 
and comparison of scalation difficult at best. 
Dorsal scale shape in Sphaerodactylus spp. 
varies from granular to elongate and strong- 
ly keeled (Fig. 8). When the scales are 
elongate they are typically extremely flat- 
tened and imbricate (Barbour, 1921). Al- 
though granular scales are common in 
related genera, truly granular scales cover- 
ing the body dorsum are rarely present in 
Sphaerodactylus (Thomas, 1975). Among 
the only exceptions are S. scapularis from 
Gorgona Island in Colombia (Harris, 1982; 
J.D.D., personal observation) and Hispan- 
iolan species such as S. cinereus, S. 
elasmorhynchus, and the extinct S. dommeli. 
Sphaerodactylus copei, another Hispaniolan 
species, has very large, swollen dorsals with 
a region of middorsal granular scales. The 
scalation of S. ciguapa may have been 
similar to this, but the fragmentary nature 
of the integument in the type makes it 
difficult to determine the actual distribution 
of scale types across the body and precise 
meristic comparisons with other congeners 
are precluded. 

Ecology. Amber inclusions have revealed a 
good deal about the floral and faunal 
composition of the Miocene biota of the 
Dominican Republic (Poinar and Poinar, 
1994, 1999). To the extent that available 
fossils permit the reconstruction of herpeto- 
faunal communities of the period, they 
appear to be similar to those predominating 
today. Species of Anolis, Sphaerodactylus, 
Typhlops, and Eleutherodactylus, the four 
genera recorded as Dominican inclusions, 
today comprise 171 of 243 species of 
amphibians and reptiles found on Hispaniola 
(Hedges, 2011). Extant Anolis lizards are 
known for their distinctive ecomorphs and 



No. 529 

have served as the basis for fruitful research 
programs in both ecology and evolution 
(Williams, 1976; Losos et al, 1998; Losos, 
2009). Hispaniolan Sphaerodactylus likewise 
seem to reflect morphological adaptation to 
particular lifestyles and substrates. Thomas 
(1975) and Thomas et al. (1992) proposed that 
head shape and coloration might be adaptive 
traits, an argument supported by replicated 
evolution of dark brown cryptic colored 
species in the montane species dwelling in the 
leaf litter from the Greater Antilles (Schwartz 
and Garrido, 1985). The study of this varia- 
tion has the potential to provide a rich system 
for the study of comparative biology (D. 
Scantlebury, personal communication). 

To date the only specimens of amber- 
preserved Sphaerodactylus that have been 
reported are S. ciguapa, two specimens of S. 
dommeli (Bohme, 1984), and an undescribed 
small (14 mm SVL + 16.6 mm tail length) 
specimen with slightly imbricated dorsal scales 
(Kluge, 1995). Kluge concluded that the last of 
these was possibly a juvenile on the basis of its 
small size, but S. ariasae, the smallest Sphaer- 
odactylus, has an adult SVL range of 14.1- 
17.9 mm (Hedges and Thomas, 2001), so the 
possibility exists that this animal represents a 
very different ecomorph from the larger S. 
ciguapa and S. dommeli. 

Although it may never be possi- 
ble to fully reconstruct the Sphaerodactylus 
fauna of the Miocene, the study of the 
relatively large number of as yet undescribed 
amber geckos from the Dominican Republic 
(Poinar and Poinar 1999; D. A. Grimaldi, 
personal communication) may result in the 
description of additional new taxa and pro- 
vide a clearer picture of the paleodiversity of 
this important group of Caribbean geckos. 


We thank James Hanken, Director of the 
Museum of Comparative Zoology, for pur- 

chasing the specimen described herein and 
making it available for our study. Yale 
Goldman provided locality data for the 
specimen and David Grimaldi (AMNH) 
provided advice on Dominican amber. Kyle 
Luckenbill and Ned Gilmore (Academy of 
Natural Sciences in Philadelphia) and San- 
dra Rareron and Kenneth Tighe (National 
Museum of Natural History, Smithsonian 
Institution) kindly helped with the X-rays. 
For access to comparative material we thank 
Richard Thomas (RT, UPRRP), Colin 
McCarthy (BMNH), Kevin de Queiroz, 
Roy McDiarmid, Ron Heyer, George Zug, 
Jeremy Jacobs, and Steve Gotte (USNM), 
Jonathan Losos and Jose Rosado (MCZ), 
Fernando Castro-H (UV-C), Hussam Zaher 
(MZUSP), Laurie Vitt (OMNH), Tony 
Gamble (JFBM), Darell Frost and David 
Kizirian (AMNH), Jens Vindum, Robert 
Drewes, and Alan Leviton (CAS), Alan 
Resetar (FMNH), and Jim McGuire and 
Carol Spencer (MVZ). Virginia Abdala and 
Esteban Lavilla (Fundacion Miguel Lillo) 
provided assistance with specimens, and Dan 
Scantlebury (University of Rochester) and 
Wolfgang Bohme and Philipp Wagner (Zo- 
ologisches Forschungsmuseum Alexander 
Koenig) kindly provided X-rays of S. shrevei 
and S. dommeli, respectively. This research 
was supported by the Gerald M. Lemole, 
M.D. Endowed Chair in Integrative Biology 
Fund, and by grant DEB 0844523 from the 
National Science Foundation of the United 


Specimens used in phylogenetic analyses 
(Sk = dry skeleton, C&S = cleared and 
stained, HRXCT = high-resolution X-ray 
computed tomography, XR = X-ray, * = 
ethanol-preserved specimens). 

Aristelliger lar (AMNH R-50272 [Sk], 
USNM 259998-260007, 260009* [XR], USNM 




260008 [C&S]), Chatogekko amazonicus 
(AMNH-R 138670 [C&S], AMNH R- 138726 
[C&S], AMNH R- 132039 [C&S], AMNH R- 
132052 [C&S], MZUSP 91394*, OMNH 36262 
[C&S], OMNH 37616 [C&S], OMNH 37274 
[C&S], OMNH 37110 [C&S], OMNH 36712 
[C&S], USNM 302283-302284*, USNM 
124173*, USNM 200660-200663*, USNM 
200664 [C&S], USNM 200665-200666*, 
USNM 288763*, USNM 288764 [C&S], 
USNM 288765-288788*, USNM 289031 
[C&S], USNM 289061-289066*, USNM 
290881-290882*, USNM 290904 [C&S], 
USNM 290944^-290945*, USNM 303472- 
303473*, USNM 570538*, USNM 304122 
304123*), Coleodactylus br achy stoma (MZUSP 
no data [C&S], MZUSP 87385*), Euleptes 
europaea (USNM 014861* [XR], USNM 
56591 1*[XR], USNM 58963 [C&S]), Gonatodes 
albogularis (AMNH R-71594[Sk], MVZ 83402 
[C&S], UV-C No data [Sk]), Gymnodactylus 
geckoides geckoides (CAS 49397 [HRXCT]), 
Hemidactylus brookii (BMNH 1978.1472*), 
Lepidoblepharis xanthostigma (RT 1875 [C&S], 
USNM 313758*, USNM 313791 [C&S], 
USNM 313834*), Narudasia f estiva (AMB 
8717 [C&S], CAS 186290 [C&S]), Phyllodacty- 
lus wirshingi (CAS 175498 [C&S], RT 13860 
[C&S]), Pristurus carted (CAS 225349 [C&S], 
BMNH 1971.44 [Sk], JFBM 15821 [Sk]), 
Pseudogekko smaragdinus (USNM 197367 
[C&S], USNM 198423 [C&S], USNM 198424 
[C&S]), Pseudogonatodes guianensis (MZUSP 
94826 [C&S], USNM 84970* [XR], USNM 
166138* [XR], USNM 234574* [XR], USNM 
316687* [XR], USNM 321059* [XR], USNM 
333018* [XR], USNM 538260-538267* [XR], 
USNM 566327* [XR]); Quedenfeldtia trachy- 
blepharus (FMNH 197682 [C&S], USNM 
71113* [XR], USNM 196417* [XR], MVZ 
178124) [C&S], Saurodactylus mauritanicus 
(BMNH [Sk], FMNH 197462 
[C&S], USNM 217454* [XR]), Sphaerodactylus 
ciguapa (MCZ R- 186380 [Sk, XR]), Sphaero- 
dactylus roosevelti (UPRRP 6376-6378 [C&S], 

UPRRP 6380-6381 [C&S], UPRRP 6488 
[C&S], USNM 326986-326987* [XR], USNM 
326996* [XR], USNM 327042* [XR]), Sphaero- 
dactylus klauberi (UPRRP 6409-6421 [C&S], 
UPRRP 6423-6427 [C&S]), Tarentola maurita- 
nica (AMNH R-71591 [Sk], AMNH R-144408 
[C&S], BMNH 1913.7.3.36 [Sk], JFBM 15824 
[Sk]), Teratoscincus scincus (BMNH 
[Sk], CAS 101437 [C&S]), Thecadactylus rapi- 
cauda (AMNH R-59722 [Sk], AMNH R-75824 
[Sk], AMNH R-85312 [Sk], BMNH 
[Sk], USNM 220204 [Sk]). 


Specimens used for comparative purposes 
(Sk = dry skeleton, C&S = cleared and 
stained, HRXCT = high-resolution X-ray 
computed tomography, XR = X-ray, * = 
ethanol-preserved specimens). 

Ailuronyx seychellensis (CAS 8421 [C&S]), 
Aristelliger georgeensis (CAS 176485 [HRX- 
CT]), Aristelliger praesignis (AMNH R-146747 
[C&S], AMNH R-71593 [Sk], AMNH R-71595 
[Sk], BMNH 1964.1812 [Sk], BMNH 
[Sk]), Coleodactylus guimaraesi (USNM 304122* 
[XR]), Coleodactylus meridionalis (MZUSP 
88673*), Coleodactylus septentrionalis (MZSP 
66554*, MZUSP 66556*, USNM 302285- 
302287* [XR], USNM 302337* [XR], USNM 
302361* [XR], USNM 531620-531622* [XR], 
USNM 566300* [XR]), Gonatodes annularis 
(USNM 535787* [XR], USNM 535791* [XR]), 
Gonatodes antillensis (AMNH R-72642 [Sk], 
USNM 94980* [XR]), Gonatodes ceciliae 
(USNM 166159* [XR]), Gonatodes humeralis 
(RT 01198 [C&S], USNM 568645* [XR]), 
USNM 568647* [XR]), USNM 568658* [XR]), 
USNM 568663* [XR]), USNM 568677* [XR]), 
USNM 568681* [XR]), USNM 568682* [XR]), 
USNM 568684* [XR]), USNM 568692* [XR]), 
Gonatodes taniae (UPRRP 006045 [C&S]), 
Lepidoblepharis buchwaldi (USNM 234565* 
[XR], USNM 234569* [XR]), Lepidoblepharis 
festae (USNM 166140-166143* [XR]), Lepido- 



No. 529 

blepharis heyerorum (USNM 217635* [XR]), 
Lepidoblepharis peraccae (UV-C 8999 [Sk]), 
Pristurus crucifer (USNM 72014* [XR], USNM 
217452* [XR], USNM 217453* [XR]), Pris- 
turus insignis (BMNH 1953.1.7.73 [Sk]), 
Pseudogonatodes barbouri (AMNH R- 
144395 [C&S], AMNH R-146746 [C&S], 
AMNH R-146752 [C&S], AMNH 146757 
[C&S]), Pseudogonatodes peruvianus (USNM 
343190* [XR], USNM 343191* [XR]), Sphaero- 
dactylus altavelensis (USNM 328548* [XR]), 
Sphaewdactylus argivus {USNM 104597* [XR]), 
Sphaerodactylus argus (USNM 251977-251978* 
[XR]), Sphaerodactylus ariasae (USNM 541804- 
541805* [XR], USNM 541807-541810* [XR]), 
Sphaerodactylus armstrongi (RT 5255 [C&S], 
USNM 260053* [XR], USNM 260046* [XR], 
USNM 260051-260054* [XR]), Sphaerodactylus 
asterulus (USNM 328946* [XR], USNM 
328949* [XR]), Sphaerodactylus beattyi (USNM 
304480-304481* [XR]), Sphaerodactylus cinereus 
(AMNH R-49566 [C&S]; USNM 292296* 
[XR]), Sphaerodactylus copei (RT 10576 [C&S], 
USNM 118881* [XR]), Sphaerodactylus corti- 
cola (USNM 211428* [XR], USNM 220548- 
220552* [XR]), Sphaerodactylus darlingtoni 
(USNM 328962* [XR]), Sphaerodactylus diffici- 
lis (AMNH R-144413-144435 [C&S], USNM 
328965* [XR]), Sphaerodactylus elegans (USNM 
27625* [XR], USNM 27981* [XR]), Sphaero- 
dactylus gfl/grae (UPRRP 6428-6432 [C&S], 
UPRRP 6434-6436 [C&S]), Sphaerodactylus 
goniorhynchus (BMNH 1963.841*), Sphaerodac- 
tylus gossei (BMNH1964.1801-2 [Sk), Sphaero- 
dactylus ladae (USNM 512248* [XR], USNM 
512251* [XR], USNM 512253-512254* [XR]), 
Sphaerodactylus leucaster (USNM 197338* 
[XR]), Sphaerodactylus levinsi (RT 8283-8284 
[Sk], USNM 220939* [XR], USNM 220921* 
[XR]), Sphaerodactylus lineolatus (BMNH*, UPRRP 3172 [C&S], USNM 
120479* [XR], USNM 120497* [XR], USNM 
12053* [XR], USNM 120504* [XR]), Sphaero- 
dactylus macrolepis (UPRRP 6437-6445 [C&S], 
USNM 221462* [XR]), Sphaerodactylus micro- 

lepis (USNM 222901* [XR]), Sphaerodactylus 
micropithecus (USNM 229891* [XR]), Sphaero- 
dactylus millepunctatus (AMNH R- 16284* 
[XR]), Sphaerodactylus monensis (UPRRP 6454 
[C&S]), Sphaerodactylus nicholsi (UPRRP 6383- 
6386 [C&S], UPRRP 63880 [C&S]), Sphaero- 
dactylus nigropunctatus (AMNH R-73470 [Sk]; 
BMNH 1946.8.24.81*), Sphaerodactylus notatus 
(BMNH 1965.186*, USNM 494822* [XR]), 
Sphaerodactylus oliveri (USNM 140431* [XR], 
USNM 140435* [XR]), Sphaerodactylus oxyrhi- 
nus (USNM 292288-292289* [XR]), Sphaero- 
dactylus pacificus (BMNH 1979.385-1979.386*, 
USNM 157531-157532* [XR]), Sphaerodactylus 
parked (USNM 328281* [XR]), Sphaerodactylus 
parthenopion (USNM 221593* [XR]), Sphaero- 
dactylus ramsdeni (USNM 309772* [XR]), 
Sphaerodactylus randi (USNM 305427-305428* 
[XR]), Sphaerodactylus rhabdotus (USNM 
292328* [XR]), Sphaerodactylus richardsonii 
(BMNH 1964.1801-2*, USNM 252126* [XR]), 
Sphaerodactylus shrevei (USNMFH 194578 
[XR]), Sphaerodactylus rosaurae (BMNH 
1946.8.16.60*, USNM 570196-570199* [XR], 
USNM 570204-570213* [XR]), Sphaerodactylus 
ruibali (USNM 78921* [XR]), Sphaerodactylus 
sabanus (USNM 27625* [XR], USNM 236098* 
[XR]), Sphaerodactylus samanensis (USNM 
319135* [XR]), Sphaerodactylus savagei (USNM 
260157* [XR]; Sphaerodactylus scapularis: 
BMNH 1901.3.29.6*, BMNH 1946.8.30.70*, 
BMNH 1902.729.1-2*, BMNH 1926.1.20*), 
Sphaerodactylus semasiops (BMNH 1968.326*, 
USNM 292294* [XR], USNM 305435* [XR]), 
Sphaerodactylus sommeri (USNM 292313* 
[XR]), Sphaerodactylus sputator (USNM 
236118* [XR]), Sphaerodactylus strep tophorus 
(USNM 541811* [XR], USNM 541813* [XR]), 
Sphaerodactylus thompsohi (USNM 328977* 
[XR]), Sphaerodactylus townsencH (UPRRP 
6389-6400 [C&S], UPRRP 6402-6407 [C&S], 
USNM 291193* [XR]), Sphaerodactylus vincenti 
(USNM 286941* [XR], USNM 121648* [XR]), 
Sphaerodactylus millepunctatus (USNM 496644* 
[XR]), Teratoscincus nucrolepis (AMNH R- 




88524 [Sk], BMNH 1934.10.9.14 [Sk]), Teratos- 
cincus przewalskii (CAS 171013 [HRXCT], 
JFBM 15826 [Sk]), Teratoscincus roboroxvskii 
(JFBM 15828 [Sk]). 


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